Association and host selectivity in multi-host pathogens.

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Association and Host Selectivity in Multi-Host Pathogens José M. Malpica1, Soledad Sacristán2, Aurora Fraile2, Fernando Garcı́a-Arenal2* 1 Departamento de Biotecnologı́a, Instituto Nacional de Investigación Agraria y Alimentaria, Madrid, Spain, 2 Departamento de Biotecnologı́a y Centro de Biotecnologı́a y Genómica de Plantas, Universidad Politécnica de Madrid, Madrid, Spain The distribution of multi-host pathogens over their host range conditions their population dynamics and structure. Also, host co-infection by different pathogens may have important consequences for the evolution of hosts and pathogens, and hostpathogen co-evolution. Hence it is of interest to know if the distribution of pathogens over their host range is random, or if there are associations between hosts and pathogens, or between pathogens sharing a host. To analyse these issues we propose indices for the observed patterns of host infection by pathogens, and for the observed patterns of co-infection, and tests to analyse if these patterns conform to randomness or reflect associations. Applying these tests to the prevalence of five plant viruses on 21 wild plant species evidenced host-virus associations: most hosts and viruses were selective for viruses and hosts, respectively. Interestingly, the more host-selective viruses were the more prevalent ones, suggesting that host specialisation is a successful strategy for multi-host pathogens. Analyses also showed that viruses tended to associate positively in co-infected hosts. The developed indices and tests provide the tools to analyse how strong and common are these associations among different groups of pathogens, which will help to understand and model the population biology of multihost pathogens. Citation: Malpica JM, Sacristán S, Fraile A, Garcı́a-Arenal F (2006) Association and Host Selectivity in Multi-Host Pathogens. PLoS ONE 1(1): e41. doi:10.1371/journal.pone.0000041 either increased or reduced virulence [e.g., 15–17]. Multiple infection of a host may also directly affect the genetic diversity of the pathogen population, as co-infection is a prerequisite for genetic exchange between different pathogen species or strains. Also, infection by one pathogen may result in an increased host susceptibility to a second pathogen, a common phenomenon named facilitation or predisposition by animal and plant pathologists, respectively [8,18]. In spite of its potential impact on pathogenicity, evolution, epidemiology and control, the distribution of pathogens over their host range and the occurrence of co-infections have been largely overlooked, and most research on pathogen ecology and epidemiology has dealt with specific pathogen-host interactions [8]. To our knowledge, it has not been analysed whether the distribution of pathogens over their host range is random or, alternatively, associations between pathogens and hosts occur, neither has been addressed whether host co-infection by different pathogens is random or associations between pathogens occur in particular hosts. Here we address these issues. First, we propose indices for the observed patterns of host infection by different pathogens, and the observed patterns of coinfection, and tests to analyse if they conform to the null hypothesis of randomness or reflect associations. Second, we apply these tests INTRODUCTION Pathogens have highly variable host ranges: in natural conditions some infect only one or a few related species (i.e., specialist pathogens) while other can infect a wide range of hosts belonging to different taxonomic groups (i.e., multi-host or generalist pathogens). A large fraction of described pathogens of humans, animals and plants are generalists [1–3]. The ability to infect different hosts conditions the epidemiology and pathogenicity of generalist pathogens and, therefore, is highly relevant for pathogen management and disease control [1,4]. The distribution of multihost pathogens over their host range, i.e. the frequency of infection in the various host species within an ecosystem, may vary largely, which could determine the population dynamics and structure of the pathogen. The distribution of a pathogen species over its host range may also determine important aspects of its biology in hosts significant from an anthropocentric viewpoint (i.e. target hosts), such as reservoirs and inoculum sources, emergence and reemergence, population thresholds for disease invasion or critical community size for disease persistence [e.g., 1,4–7]. Animal or plant species may be hosts for a range of pathogens, and most host populations encounter a large number of different pathogen species [8]. For significant host species, there is abundant evidence of differences in the infection frequency of the various pathogen species present in an ecosystem. The distribution of pathogens over their hosts, and the distribution of different pathogens within a host species, will affect the frequency of multiple infection of an individual host by different pathogens. Multiple infection may have important consequences for the infected hosts, for the pathogens, and for host-pathogen coevolution [8,9]. In the host, frequent co-infections may lead to heterozygote superiority against multiple pathogens and contribute to the persistence in host populations of alleles conferring susceptibility to disease [10]. In multiple infected hosts, pathogens can cooperate or can compete for host resources, which will affect each other’s fitness. Hence, multiple infections will be a factor in pathogen evolution. Theoretical analyses predict that the withinhost dynamics of microparasites in multiple infected hosts may have important consequences in the evolution of their virulence [11–14], and there is evidence that multiple infection may result in PLoS ONE | Academic Editor: Angus Buckling, Oxford University, United Kingdom Received August 31, 2006; Accepted October 2, 2006; Published December 20, 2006 Copyright: ß 2006 Malpica et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This work was in part supported by grant AGL2004-03496, Ministerio de Educación y Ciencia, Spain, to AF. SS was in receipt of a FPU fellowship, Ministerio de Educación y Ciencia, Spain. Competing Interests: The authors have declared that no competing interests exist. * To whom correspondence should be addressed. E-mail: 1 December 2006 | Issue 1 | e41 Associations in Pathogens 102 plants were infected by at least one virus out of 1060 analysed plants (Table 4). The above described test of association between pathogens was applied to this set. The data in Table 4 showed a tendency of the analysed viruses to associate positively: the distribution of the association index (AI) was skewed towards positive values (Fig. 3) so that out of 68 AIs computed for the five viruses in 16 plant species, 47/68 (more than two thirds) were positive and 21/68 were negative. Moreover, there was a conspicuous tendency of the positive AI values to have smaller probabilities (r = 20.6575, P,1024, in a Spearman rank correlation test). When the pooled sample from the sixteen plant species was considered, the AI was positive and significantly different from zero for each of the five viruses, i.e. each of the five viruses was found in co-infection with a frequency significantly higher than expected from the null hypothesis of independence of infection. However, this was not so when the data for each of the sixteen plant species were analyzed separately. Hence the association analysis uncovered two patterns that were not obvious: i) a general tendency of the analysed viruses to associate positively, ii) association depended on both the plant and the virus species. to data on the prevalence of five insect-borne virus species in wild plant species within an agroecosystem in Central Spain. Results of these analyses uncover patterns that, if general, would be highly relevant to understand the ecology and evolution of pathogens. RESULTS Association between viruses and hosts, and among viruses, in weeds in Central Spain Between January 2000 and December 2002, 2275 samples from 56 weed species or genera pertaining to 21 dicotyledonous plant families, were collected and analyzed for infection by the aphid transmitted viruses Alfalfa mosaic virus (AMV, genus Alfamovirus, family Bromoviridae), Beet western yellows virus (BWYV, genus Polerovirus, family Luteoviridae), Cucumber mosaic virus (CMV, genus Cucumovirus, family Bromoviridae), and Watermelon mosaic virus (WMV, genus Potyvirus, family Potyviridae), and by the thrips-transmitted Tomato spotted wilt virus (TSWV, genus Tospovirus, family Bunyaviridae) [19]. Except for TSWV, which has a single-stranded RNA genome of negative and ambisense polarity, all other viruses have single-stranded RNA genomes of messenger polarity. AMV, CMV and WMV are transmitted by aphids in a non-persistent manner, i.e. the virus is retained in the distal structures of the aphid mouth parts for short period of time. BWYV is transmitted in a circulative, non-propagative manner, i.e., the virus penetrates through the gut wall into the haemocoel of the insect vector, and circulates with the haemolymph to reach the salivary glands, from where it is inoculated into new plants. TSWV follows a similar path within the thrips body, but infects and multiplies in the insect cells [19]. All five viruses cause important diseases in vegetable crops worldwide, including the studied region in Central Spain, but infection in the analysed wild hosts was asymptomatic. Table 1 shows the number of samples analysed and the number of infected plants by each of these five virus species, in single or multiple infection, in the 21 most frequently found plant species in three monitored habitats (see Methods) for the analysed period. To this data set tests for association between hosts and pathogens (see Methods) were applied. The index of selectivity of pathogen (ISP), and its significance, is shown in Table 2 for the five viruses. The distribution of three of five analysed viruses over their hosts was significantly non-random, i.e. some of the available hosts were preferentially infected. Fig. 1 shows the relationship between prevalence and the ISP for the five viruses. A positive correlation was found for both parameters (r = 0.9347, P = 0.0189 in a Spearman rank correlation test), i.e., the more host-selective viruses were those with a highest prevalence in the analysed ecosystem. Similarly, the index of selectivity of the host (ISH), and its significance, was calculated for the 21 host plant species in Table 1, and values are shown in Table 3. For about half (9/21) of the analysed hosts (Amaranthus spp., Cirsium arvense, Convolvulus arvensis, Diplotaxis erucoides, Lactuca serriola, Medicago sativa, Portulaca oleracea, Solanum nigrum and Taraxacum spp.) differences in the prevalence of the five viruses departed significantly from random. Fig. 2 shows the relationship between virus prevalence and the ISH for the 21 host species. Again, a positive correlation between both parameters was found (r = 0.5161, P = 0.0166, in a Spearman rank correlation test), i.e., the more virus-selective hosts were those with a higher prevalence of virus infection. The relationships between prevalence and selectivity for viruses and hosts were not due to a coincidence in the frequency of infection among hosts by different viruses, as shown by a contingency analysis of counts of infected hosts by the different viruses (P,1024). For 16 of the 21 plant species in Table 1, co-infection with more than one of the five viruses occurred. For these 16 plant species, PLoS ONE | DISCUSSION Most efforts to understand the population biology of pathogens have focussed on specialist pathogens, and population biologists have successfully developed a formal understanding of the dynamics and evolution of single-host pathogens. However, most pathogens of humans, animals and plants are multi-host pathogens [1–3,20]. As stated by Woolhouse et al. [1] ‘‘understanding the more complex population biology of multi-host pathogens will be one major challenge in the 21st century ‘‘. There is evidence that within an ecosystem the prevalence of multi-host pathogens may differ largely for the different species of their host range [e.g., [21– 25]]. Similarly, there is evidence of large differences in the prevalence on a host species of the various pathogens that are able to infect it [e.g., [26–29]]. However, no attempt has been made, to our knowledge, to analyse if differences in the distribution of multihost pathogens over their hosts are random or if there are associations between hosts and pathogens. The uncovering of associations between hosts and pathogens would be highly relevant to understand and model the population biology of multi-host pathogens, and for understanding the phenomenon of generalism itself. We present here indices and tests to analyse if there is association between multi-host pathogens and their hosts. The proposed indices of selectivity for the pathogen and for the host measure the degree of association between hosts and pathogens. The tests analyse the homogeneity of distribution of a pathogen over different host species or populations, and of different pathogens on a host, and analyse how significantly the values of the indices departs from zero (i.e. no association). The literature on pathogen ecology does not abound with data on the prevalence of various pathogens on various hosts. Hence, we have applied these indices to our unpublished data on the prevalence of five insectborne plant viruses on 21 species of wild plants in an agroecosystem in central Spain over a three year period. The analysis of the prevalence of the different viruses in each host species by the homogeneity test that we propose, shows that half of the analysed plant species showed an index of selectivity of the host (ISH) significantly different from zero. The distribution of the host species showing virus selectivity was not related to taxonomy, habitat (fallow fields, edges or wastelands), seasonality or vegetative cycle (annual vs. perennial) (not shown). Interestingly, there was a positive correlation between the ISH and the average virus prevalence for these 21 host plant species, showing that the 2 December 2006 | Issue 1 | e41 3 Multiple infections3 AMV+ Single infections 1 2 AMV BWYV CMV TSWV WMV Amaranthus spp. 87 1 5 31 3 4 C. bursa-pastoris (L.) Medicus 5 0 0 0 1 0 Chenopodium album L. 69 4 0 3 1 1 Cirsium arvense (L.) Scop. 42 8 8 20 1 3 Convolvulus arvensis L. 113 10 6 7 0 0 Conyza sp. 121 5 10 7 5 1 Datura stramonium L. 26 4 2 2 0 3 2 Diplotaxis erucoides (L.) DC. 164 15 11 44 11 2 1 Host species N Lactuca serriola L. 66 1 1 13 1 1 Lamium amplexicaule L. 35 2 2 5 2 0 Malva sylvestris L. 4 0 0 1 0 0 Medicago sativa L. 90 38 4 9 5 5 Papaver rhoeas L. 61 1 0 0 3 0 B C BMV+ T W B+C B+T C+T T+W B+C+T B+T+W 1 C CMV+ T C+T C+T+W B+T 3 B+W 3 1 4 1 3 5 1 1 1 1 2 1 2 2 1 3 1 1 1 2 1 3 1 2 3 1 1 1 1 1 3 1 2 Plantago sp 61 7 6 7 4 0 Portulaca oleracea L. 7 0 0 5 0 0 Silybum marianum (L.)Gaertner 29 1 2 1 1 0 Solanum nigrum L. 24 0 0 4 0 0 Sonchus oleraceus L. 51 3 1 4 3 2 Taraxacum sp. 15 0 0 10 0 0 Trifolium pratense L. 37 1 0 2 1 1 1 Xanthium strumarium Moretti. 8 1 0 1 1 1 1 1 1 1 1 1 Amaranthus spp: A. retroflexus (L.) or A. hybridus (L.); C. bursa-pastoris (L.) Medicus: Capsella bursa-pastoris (L.) Medicus; Conyza spp: C. canadensis (L.) Cronq. or C. bonaerenses (L.) Cronq.; Plantago spp: P. coronopus L. or P. lagopus L.; Taraxacum spp: undetermined species. N = Number of analysed plants 3 A = AMV, B = BWYV, C = CMV, T = TSWV, W = WMV doi:10.1371/journal.pone.0000041.t001 2 Associations in Pathogens December 2006 | Issue 1 | e41 . PLoS ONE | Table 1. Number of single and multiple infections with AMV, BWYV, CMV, TSWV and WMV in twenty one weed species . . . Associations in Pathogens Table 3. Average prevalence of virus infection (Pk), and index of selectivity of the host (ISH) for twenty one weed species. . Table 2. Average prevalence (Pi), and index of selectivity of the pathogen (ISP) for five virus species. . Species Pi ISH P1 AMV 0.128 0.3943 0.000 BWYV 0.065 0.2193 0.000 CMV 0.173 0.3838 0.000 TSWV 0.039 0.1398 0.359 WMV 0.022 0.1844 0.063 1 Probability of rejection of the null hypothesis of homogeneity of the prevalence of five analysed viruses. Raw significance probabilities were corrected by the sequential Bonferroni method for multiple independent tests as indicated in [60]. doi:10.1371/journal.pone.0000041.t002 more selective hosts are more prone to be virus-infected, obviously by the virus(es) that better infects them. This phenomenon suggests that in spite that each host encounters a wide array of pathogens, mechanisms of escape and/or resistance [30] to some of equilibrium value of MeCpG steps (,+14 deg.) [31,44]. In comparison, methylation has a significantly lower stability cost when happening at major groove positions, such as 211 and 21 base pair from dyad (mutations 9 and 12), where the roll of the nucleosome bound conformation (+10 deg.) is more compatible with the equilibrium geometry of MeCpG steps. The nucleosome destabilizing effect of cytosine methylation increases with the number of methylated cytosines, following the same position dependence as the single methylations. The multiple-methylation case reveals that each major groove meth- PLOS Computational Biology | 3 November 2013 | Volume 9 | Issue 11 | e1003354 DNA Methylation and Nucleosome Positioning ylation destabilizes the nucleosome by around 1 kJ/mol (close to the average estimate of 2 kJ/mol obtained for from individual methylation studies), while each minor groove methylation destabilizes it by up to 5 kJ/mol (average free energy as single mutation is around 6 kJ/mol). This energetic position-dependence is the reverse of what was observed in a recent FRET/SAXS study [30]. The differences can be attributed to the use of different ionic conditions and different sequences: a modified Widom-601 sequence of 157 bp, which already contains multiple CpG steps in mixed orientations, and which could assume different positioning due to the introduction of new CpG steps and by effect of the methylation. The analysis of our trajectories reveals a larger root mean square deviation (RMSD) and fluctuation (RMSF; see Figures S2– S3 in Text S1) for the methylated nucleosomes, but failed to detect any systematic change in DNA geometry or in intermolecular DNA-histone energy related to methylation (Fig. S1B, S1C, S4–S6 in Text S1). The hydrophobic effect should favor orientation of the methyl group out from the solvent but this effect alone is not likely to justify the positional dependent stability changes in Figure 2, as the differential solvation of the methyl groups in the bound and unbound states is only in the order of a fraction of a water molecule (Figure S5 in Text S1). We find however, a reasonable correlation between methylation-induced changes in hydrogen bond and stacking interactions of the bases and the change in nucleosome stability (see Figure S6 in Text S1). This finding suggests that methylation-induced nucleosome destabilization is related to the poorer ability of methylated DNA to fit into the required conformation for DNA in a nucleosome. Changes in the elastic deformation energy between methylated and un-methylated DNA correlate with nucleosomal differential binding free energies To further analyze the idea that methylation-induced nucleosome destabilization is connected to a worse fit of methylated DNA into the required nucleosome-bound conformation, we computed the elastic energy of the nucleosomal DNA using a harmonic deformation method [36,37,44]. This method provides a rough estimate of the energy required to deform a DNA fiber to adopt the super helical conformation in the nucleosome (full details in Suppl. Information Text S1). As shown in Figure 2, there is an evident correlation between the increase that methylation produces in the elastic deformation energy (DDE def.) and the free energy variation (DDG bind.) computed from MD/TI calculations. Clearly, methylation increases the stiffness of the CpG step [31], raising the energy cost required to wrap DNA around the histone octamers. This extra energy cost will be smaller in regions of high positive roll (naked DNA MeCpG steps have a higher roll than CpG steps [31]) than in regions of high negative roll. Thus, simple elastic considerations explain why methylation is better tolerated when the DNA faces the histones through the major groove (where positive roll is required) that when it faces histones through the minor groove (where negative roll is required). Nucleosome methylation can give rise to nucleosome repositioning We have established that methylation affects the wrapping of DNA in nucleosomes, but how does this translate into chromatin structure? As noted above, accumulation of minor groove methylations strongly destabilizes the nucleosome, and could trigger nucleosome unfolding, or notable changes in positioning or phasing of DNA around the histone core. While accumulation of methylations might be well tolerated if placed in favorable positions, accumulation in unfavorable positions would destabilize the nucleosome, which might trigger changes in chromatin structure. Chromatin could in fact react in two different ways in response to significant levels of methylation in unfavorable positions: i) the DNA could either detach from the histone core, leading to nucleosome eviction or nucleosome repositioning, or ii) the DNA could rotate around the histone core, changing its phase to place MeCpG steps in favorable positions. Both effects are anticipated to alter DNA accessibility and impact gene expression regulation. The sub-microsecond time scale of our MD trajectories of methylated DNAs bound to nucleosomes is not large enough to capture these effects, but clear trends are visible in cases of multiple mutations occurring in unfavorable positions, where unmethylated and methylated DNA sequences are out of phase by around 28 degrees (Figure S7 in Text S1). Due to this repositioning, large or small, DNA could move and the nucleosome structure could assume a more compact and distorted conformation, as detected by Lee and Lee [29], or a slightly open conformation as found in Jimenez-Useche et al. [30]. Using the harmonic deformation method, we additionally predicted the change in stability induced by cytosine methylation for millions of different nucleosomal DNA sequences. Consistently with our calculations, we used two extreme scenarios to prepare our DNA sequences (see Fig. 3): i) all positions where the minor grooves contact the histone core are occupied by CpG steps, and ii) all positions where the major grooves contact the histone core are occupied by CpG steps. We then computed the elastic energy required to wrap the DNA around the histone proteins in unmethylated and methylated states, and, as expected, observed that methylation disfavors DNA wrapping (Figure 3A). We have rescaled the elastic energy differences with a factor of 0.23 to match the DDG prediction in figure 2B. In agreement with the rest of our results, our analysis confirms that the effect of methylation is position-dependent. In fact, the overall difference between the two extreme methylation scenarios (all-in-minor vs all-in-major) is larger than 60 kJ/mol, the average difference being around 15 kJ/ mol. We have also computed the elastic energy differences for a million sequences with CpG/MeCpG steps positioned at all possible intermediate locations with respect to the position (figure 3B). The large differences between the extreme cases can induce rotations of DNA around the histone core, shifting its phase to allow the placement of the methylated CpG steps facing the histones through the major groove. It is illustrative to compare the magnitude of CpG methylation penalty with sequence dependent differences. Since there are roughly 1.5e88 possible 147 base pairs long sequence combinations (i.e., (4n+4(n/2))/2, n = 147), it is unfeasible to calculate all the possible sequence effects. However, using our elastic model we can provide a range of values based on a reasonably large number of samples. If we consider all possible nucleosomal sequences in the yeast genome (around 12 Mbp), the energy difference between the best and the worst sequence that could form a nucleosome is 0.7 kj/mol per base (a minimum of 1 kJ/mol and maximum of around 1.7 kJ/mol per base, the first best and the last worst sequences are displayed in Table S3 in Text S1). We repeated the same calculation for one million random sequences and we obtained equivalent results. Placing one CpG step every helical turn gives an average energetic difference between minor groove and major groove methylation of 15 kJ/ mol, which translates into ,0.5 kJ/mol per methyl group, 2 kJ/ mol per base for the largest effects. Considering that not all nucleosome base pair steps are likely to be CpG steps, we can conclude that the balance between the destabilization due to CpG methylation and sequence repositioning will depend on the PLOS Computational Biology | 4 November 2013 | Volume 9 | Issue 11 | e1003354 DNA Methylation and Nucleosome Positioning Figure 3. Methylated and non-methylated DNA elastic deformation energies. (A) Distribution of deformation energies for 147 bplong random DNA sequences with CpG steps positioned every 10 base steps (one helical turn) in minor (red and dark red) and major (light and dark blue) grooves respectively. The energy values were rescaled by the slope of a best-fit straight line of figure 2, which is 0.23, to por la lectura a través de la lectura de la prensa. La educación en los medios las fuerzas dispersas en función de los soportes mediáticos y orientarse más hacia la educación en medios que al dominio adquiere pleno derecho y entidad en la sección sexta titulada «competencias sociales y cívi- técnico de los aparatos. cas» que indica que «los alum- nos deberán ser capaces de juz- gar y tendrán espíritu crítico, lo que supone ser educados en los las programaciones oficiales, ya que, a lo largo de un medios y tener conciencia de su lugar y de su influencia estudio de los textos, los documentalistas del CLEMI en la sociedad». han podido señalar más de una centena de referencias a la educación de los medios en el seno de disciplinas 4. Un entorno positivo como el francés, la historia, la geografía, las lenguas, Si nos atenemos a las cifras, el panorama de la las artes plásticas : trabajos sobre las portadas de educación en medios es muy positivo. Una gran ope- prensa, reflexiones sobre temas mediáticos, análisis de ración de visibilidad como la «Semana de la prensa y publicidad, análisis de imágenes desde todos los ángu- de los medios en la escuela», coordinada por el CLE- los, reflexión sobre las noticias en los países europeos, MI, confirma año tras año, después de 17 convocato- información y opinión rias, el atractivo que ejerce sobre los profesores y los Esta presencia se constata desde la escuela mater- alumnos. Concebida como una gran operación de nal (2 a 6 años) donde, por ejemplo, se le pregunta a complementariedad entre la escuela y los profesiona- los niños más pequeños si saben diferenciar entre un les de los medios, alrededor del aprendizaje ciudada- periódico, un libro, un catálogo, a través de activida- no de la comunicación mediática, este evento moviliza des sensoriales, si saben para qué sirve un cartel, un durante toda una semana un porcentaje elevado de periódico, un cuaderno, un ordenador si son capa- centros escolares que representan un potencial de 4,3 ces de reconocer y distinguir imágenes de origen y de millones de alumnos (cifras de 2006). Basada en el naturaleza distintas. Podríamos continuar con más voluntariado, la semana permite desarrollar activida- ejemplos en todos los niveles de enseñanza y práctica- des más o menos ambiciosas centradas en la introduc- Páginas 43-48 ción de los medios en la vida de la escuela a través de la instalación de kioscos, organización de debates con profesionales y la confección por parte de los alumnos de documentos difundidos en los medios profesionales. Es la ocasión de dar un empujón a la educación en medios y de disfrutarlos. Los medios –un millar en 2006– se asocian de maneras diversas ofreciendo ejemplares de periódicos, acceso a noticias o a imágenes, proponiendo encuentros, permitiendo intervenir a los jóvenes en sus ondas o en sus columnas Esta operación da luz al trabajo de la educación en medios y moviliza a los diferentes participantes en el proyecto. 5. La formación de los docentes La formación es uno de los pilares principales de la educación en los medios. Su función es indispensable ya que no se trata de una disciplina, sino de una enseñanza que se hace sobre la base del voluntariado y del compromiso personal. Se trata de convencer, de mostrar, de interactuar. En primer lugar es necesario incluirla en la formación continua de los docentes, cuyo volumen se ha incrementado desde 1981 con la aparición de una verdadera política de formación continua de personal. Es difícil dar una imagen completa del volumen y del público, pero si nos atenemos a las cifras del CLEMI, hay más de 24.000 profesores que han asistido y se han involucrado durante 2004-05. 5.1. La formación continua En la mayoría de los casos, los profesores reciben su formación en contextos cercanos a su centro de trabajo, o incluso en este mismo. Después de una política centrada en la oferta que hacían los formadores, se valora más positivamente la demanda por parte del profesorado, ya que sólo así será verdaderamente fructífera. Los cursos de formación se repartieron en varias categorías: desde los formatos más tradicionales (cursos, debates, animaciones), hasta actividades de asesoramiento y de acompañamiento, y por supuesto los coloquios que permiten un trabajo en profundidad ya que van acompañados de expertos investigadores y profesionales. Citemos, por ejemplo en 2005, los coloquios del CLEMI-Toulouse sobre el cine documental o el del CLEMI-Dijon sobre «Políticos y medios: ¿connivencia?». Estos coloquios, que forman parte de un trabajo pedagógico regular, reagrupan a los diferentes participantes regionales y nacionales alrededor de grandes temas de la educación en medios y permiten generar nuevos conocimientos de aproximación y una profundización. Páginas 43-48 Hay otro tipo de formación original que se viene desarrollando desde hace menos tiempo, a través de cursos profesionales, como por ejemplo, en el Festival Internacional de Foto-periodismo «Visa para la imagen», en Perpignan. La formación se consolida en el curso, da acceso a las exposiciones, a las conferencias de profesionales y a los grandes debates, pero añade además propuestas pedagógicas y reflexiones didácticas destinadas a los docentes. Estas nuevas modalidades de formación son también consecuencia del agotamiento de la formación tradicional en las regiones. Los contenidos más frecuentes en formación continua conciernen tanto a los temas más clásicos como a los cambios que se están llevando a cabo en las prácticas mediáticas. Así encontramos distintas tendencias para 2004-05: La imagen desde el ángulo de la producción de imágenes animadas, el análisis de la imagen de la información o las imágenes del J.T. La prensa escrita y el periódico escolar. Internet y la información en línea. Medios y educación de los medios. 5.2 La formación inicial La formación inicial está aun en un grado muy ini- cial. El hecho de que la educación en medios no sea una disciplina impide su presencia en los IUFM (Institutos Universitarios de Formación de Maestros) que dan una prioridad absoluta a la didáctica de las disciplinas. En 2003, alrededor de 1.400 cursillistas sobre un total de 30.000 participaron en un momento u otro de un módulo de educación en medios. Estos módulos se ofrecen en función del interés que ese formador encuentra puntualmente y forman parte a menudo de varias disciplinas: documentación, letras, historia-geografía Estamos aún lejos de una política concertada en este dominio. La optativa «Cine-audiovisual» ha entrado desde hace muy poco tiempo en algunos IUFM destinada a obtener un certificado de enseñanza de la opción audiovisual y cine. Internet tiene cabida también en los cursos de formación inicial, recientemente con la aparición de un certificado informático y de Internet para los docentes, dirigido más a constatar competencias personales que a valorar una aptitud para enseñarlos. 6. ¿Y el futuro? El problema del futuro se plantea una vez más por la irrupción de nuevas técnicas y nuevos soportes. La difusión acelerada de lo digital replantea hoy muchas cuestiones relativas a prácticas mediáticas. Muchos Comunicar, 28, 2007 47 Comunicar, 28, 2007 Enrique Martínez-Salanova '2007 para Comunicar 48 trabajos que llevan el rótulo de la educación en medios solicitan una revisión ya que los conceptos cambian. La metodología elaborada en el marco de la educación en medios parece incluso permitir la inclinación de la sociedad de la información hacia una sociedad del conocimiento, como defiende la UNESCO. En Francia, se necesitaría unir las fuerzas dispersas en función de los soportes mediáticos y orientarse más hacia la educación en medios que al dominio técnico de los aparatos. Los avances recientes en el reconocimiento de estos contenidos y las competencias que supondrían podrían permitirlo. Referencias CLEMI/ACADEMIE DE BORDEAUX (Ed.) (2003): Parcours médias au collège: approches disciplinaires et transdisciplinaires. Aquitaine, Sceren-CRDP. GONNET, J. (2001): Education aux médias. Les controverses fécondes. Paris, Hachette Education/CNDP. SAVINO, J.; MARMIESSE, C. et BENSA, F. (2005): L’éducation aux médias de la maternelle au lycée. Direction de l’Enseignement Scolaire. Paris, Ministère de l’Education Nationale, Sceren/CNDP, Témoigner. BEVORT, E. et FREMONT, P. (2001): Médias, violence et education. Paris, CNDP, Actes et rapports pour l’éducation. – fiches pédagogiques, rapports et liens avec les pages régionales/académiques. – Le site «Quai des images» est dédié à l’enseignement du cinéma et de l’audiovisuel. – la rubrique «Côté profs» a une entrée «education aux médias». – Programme européen d’éducation aux risques liés à Internet. dResedfeleexliobnuetsacón Páginas 43-48
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